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Parajulid milliped studies XI : Initial assessment of the tribe Gosiulini (Diplopoda: Julida)
(2016)
The parajulid milliped tribe Gosiulini (Diplopoda: Julida) comprises two genera – Gosiulus Chamberlin, with three projections on the posterior gonopod and two species in the southcentral/southwestern United States (US) [Arizona, Colorado, New Mexico, and Texas], and monotypic Minutissimiulus Shelley, n. gen., with two projections, in Nuevo León, Mexico. Gosiulus conformatus Chamberlin occupies the plains/fl atlands of Texas, while its congener inhabits high elevations to the west in all four US states. Both are anticipated in Mexico (Coahuila, Chihuahua, and Sonora), and G. conformatus is expected in southeastern Colorado, eastern New Mexico, and the Oklahoma panhandle. The eastern boundary of G. conformatus and the genus/tribe conforms to the western border of the Piney Woods biome in eastern Texas. As shown by the posterior gonopod drawing in the original description, Parajulus timpius Chamberlin, previously considered of “uncertain generic position or validity,” is unquestionably the oldest name for the western species. The anteriormost posterior gonopod projection, absent from Minutissimiulus, is considered the “prefemoral process,” while the “solenomere” and a third branch arise from a common base.
Because of positional homology with “process ‘C’” in Nesoressini, the last projection is accorded this name, which may also apply to the “prefemoral process” in Aniulini. Minutissimiulus biramus Shelley, n. sp., is proposed along with the following new subjective synonymies: Apacheiulus Loomis under Gosiulus; Ziniulus aethes and Z. medicolens, both by Chamberlin, and Z. ambiguus and Z. nati, both by Loomis, under G. conformatus; and A. pinalensis and A. guadelupensis, both by Loomis, under G. timpius, new combination. Ziniulus navajo Chamberlin becomes an objective synonym of P. timpius because its holotype is designated neotype of the latter. Minutissimiulus biramus Shelley is the fi rst Mexican gosiuline and “mainland” Mexican parajulid not in the tribe Parajulini.
Hiltonius carpinus carpinus Chamberlin, 1943 (Spirobolida: Spirobolidae), is authoritatively recorded from the United States for the first time; it is known only from southern/southeastern Arizona but should be expected in adjoining counties of New Mexico. The northernmost locality is the Pinaleno Mountains, Graham County, and its distribution extends to southern Mexico; the other subspecies, H. c. vulcan (Chamberlin, 1953), occurs in Guatemala. The range of H. c. carpinus includes the type locality of the enigmatic H. fossulifer (Pocock, 1908), lending credence to prior suggestions that the names are synonymous. Three new Mexican states – Durango, Jalisco, and Nuevo León – are documented for H. c. carpinus.
The diplopod orders Callipodida and Polydesmida, and their respective families Abacionidae and
Xystodesmidae, are initially recorded from South Dakota as is Polydesmidae from North Dakota. Other new records of
indigenous taxa include Abacion Rafinesque, 1820/A. texense (Loomis, 1937) and Pleuroloma/P. flavipes, both by
Rafinesque, 1820, from South Dakota, and Pseudopolydesmus Attems, 1898/P. serratus (Say, 1821) from Alabama,
Connecticut, Delaware, New Hampshire, North Dakota, South Carolina, and the District of Columbia. New records of
Aniulus garius Chamberlin, 1912, A. (Hakiulus) d. diversifrons (Wood, 1867), and Oriulus venustus (Wood, 1864)
(Julida: Parajulidae) are provided for western Minnesota and/or eastern North Dakota. Published records from these
states are summarized, and the introduced taxa, Julidae/Cylindroiulus Verhoeff, 1894/C. caeruleocinctus (Wood, 1864)
and Paradoxosomatidae/Oxidus Cook, 1911/O. gracilis (C. L. Koch, 1847), are newly recorded from the Dakotas. The
distribution of P. serratus, which extends from Maine to South Carolina and the Florida panhandle, west to Texas, and
north to Fargo, North Dakota is described and discussed. This distribution exhibits a prominent southeastern lacuna
which we hypothesize suggests replacement by younger, more successful species, as postulated for a similar distributional
gap in Scytonotus granulatus (Say, 1821).
The endemic Floridian milliped genus, Floridobolus Causey, 1957, more closely related to tylobolinines in the western United States (US), Mexico, and Guatemala than syntopic spirobolines, is incorporated into Spirobolidae (Spirobolida: Spirobolidea). With taxonomic priority by one year, its monotypic family is reduced to Floridobolinae, n. stat., comprising Floridobolini and Tylobolini, n. stats., the counterpart to Spirobolinae, comprising Spirobolini and Aztecolini, n. tribe; relationships are Floridobolini + (Tylobolini + (Aztecolini + Spirobolini)). Like F. penneri Causey, 1957, 208 km (130 mi) to the south in the Lake Wales Ridge, Polk and Highlands counties (cos.), F. orini n. sp., inhabits “Big Scrub” environments in the Ocala National Forest, Marion Co. Biogeographic reconstructions, compatible with broader hypotheses on the class’ evolutionary history, indicate that, from a presumptive source area in northern Mexico where the subfamilies overlap, spirobolid stock penetrated the “proto-US” four times, once per tribe, before the Western Interior Seaway developed in the Cretaceous Period, Mesozoic Era. Three expansions headed northeastward into future “Appalachia,” from which taxa spread southward as the Seaway receded. Floridobolini, the fi rst invader, had to be in “proto-Georgia” and positioned to penetrate Florida when the sand dunes that comprise the “Central Highlands” emerged from the sea in the Oligocene (Cenozoic), ~25 mya. As sea levels rose and fell, the dunes fragmented into islands and the subcontinuous Floridobolus population was partitioned. The southernmost became F. penneri; F. orini inhabited a northern island; and a graduate student is investigating other insular remnants for additional species. Shortly after Floridobolini began spreading, Hiltonius/ Tylobolini arose and expanded both southward to Guatemala and northwestward to California; Tylobolus Cook, 1904, diverged in the latter area and dispersed northward to Washington and eastward to Utah/Arizona. The third invader, and the second to disperse northeastward, was Aztecolini, which probably eradicated Floridobolini from some of its established range and was partitioned into Mexican (Aztecolus Chamberlin, 1943) and US (Chicobolus Chamberlin, 1947) taxa by the Seaway. The fi nal invader, Spirobolini, dispersed northwestward and northeastward to both the Pacifi c and Atlantic coasts; instead of Trans-Beringia, we prefer penetration of the Asian part of “Asiamerica,” when it temporarily formed during the Cretaceous, to explain the Mongolian fossil genus, Gobiulus Dzik, 1975, herein assigned to Tylobolini, and the occurrence of Spirobolus Brandt, 1833, in China and Taiwan today. In the east, Narceus Rafi nesque, 1820, spread across Appalachia, eradicated most remaining populations of Floridobolus and Chicobolus, and expanded to Maine and Québec after retreat of the Wisconsin glaciation. Chicobolus and Narceus also penetrated earliest Florida; the former established itself in the Central Highlands, spread through the widening peninsula as sea levels fell, and remained on insular refugia when waters rose. Apparently fueled by the different Floridian environments, Narceus underwent time-consuming speciation; consequently, Floridobolus and Chicobolus still survive on the peninsula, and an allopatric population of the latter inhabits coastal South Carolina. However, N. gordanus (Chamberlin, 1943) occurs syntopically with both in peninsular Florida and may be actively eradicating them from their last stronghold. Trigoniulus niger, takahasii, and segmentatus, all by Takakuwa, 1940, are removed from Spirobolidae and returned toTrigoniulidae (Trigoniulidea). New records in the Appendix include the fi rst of Aztecolus from Durango and Jalisco, Mexico.
The southern Appalachian millipeds Boraria stricta (Brölemann, 1896) and B. infesta (Chamberlin, 1918) (Diplopoda: Polydesmida: Xystodesmidae) have become established in Westchester Co., New York, and Hartford Co., Connecticut, respectively. Only three individuals are available for the latter, but B. stricta has established a reproducing population in southern New York state. This species is also recorded from Bland Co., Virginia, in the Ridge and Valley Physiographic Province. Boraria profuga (Causey, 1955) comprises two allopatric populations, one in Montgomery Co., Arkansas, and the other in Ouachita Parish, Louisiana. Distributional records and gonopod drawings are presented for these species plus B. deturkiana (Causey, 1942).
Localities are documented for the milliped Abacion texense (Loomis, 1837) (Callipodida: Abacionidae) whose distribution forms both the northern and southern ordinal limits in the Western Hemisphere. The westernmost component of Abacion Rafinesque, 1820, A. texense is the only milliped species whose range spans the Mississippi and Pecos rivers and the Rio Grande. Distribution extremes are in Hennepin County (Co.), Minnesota, in the north; Terrell and Potter cos., Texas, in the west; Alcorn Co., Mississippi, in the east; and southwestern Tamaulipas, Mexico, in the south. Occurrences are projected for southeastern South Dakota, northwestern Alabama, and the southwestern periphery of Tennessee. The type series of A. texense consists solely of the male holotype, so a neotype will be needed if this individual is ever lost, because no paratypes were officially designated.
The chilopod, Cryptops hortensis (Donovan, 1810) (Scolopendromorpha: Cryptopidae), and the diplopods, Pseudospirobolellus avernus (Butler, 1876) (Spirobolida: Pseudospirobolellidae) and Oxidus gracilis (C. L. Koch, 1847) (Polydesmida: Paradoxosomatidae), are newly recorded from Saba Island, Lesser Antilles, which also harbors one additional scolopendromorph and four more chilognath millipeds. Except for the plausibly native scolopendrid centipede, Scolopendra alternans Leach, 1813, all are human introductions. Concentrated sampling is needed in the cloud/elfin forest atop Mt. Scenery, where indigenous millipeds may reside, and with extraction techniques throughout the island, to potentially document the diplopod subclass Penicillata. Nine small Caribbean islands in addition to Saba have been incorrectly reported as lacking diplopod records because publications citing them were overlooked by past authors. Works documenting myriapods from small Caribbean islands are consolidated.
The biogeographic significance of Diplopoda is substantiated by 50 maps documenting indigenous occurrences of the 16 orders, the three Spirostreptida s. l. suborders – Cambalidea, Epinannolenidea, Spirostreptidea – and all higher taxa including Diplopoda itself. The class is indigenous to all continents except Antarctica and islands/archipelagos in all temperate and tropical seas and oceans except the Arctic; it ranges from Kodiak Island and the northern Alaskan Panhandle, United States (USA), southern Hudson Bay, Canada, and near or north of the Arctic Circle in Iceland, continental Scandinavia, and Siberia to southern “mainland” Argentina, the southern tips of Africa and Tasmania, and Campbell Island, subantarctic New Zealand. The vast, global distribution is interrupted by sizeable, poorly- or unsampled areas including the Great Basin, USA; the Atacama Desert region of Chile and neighboring countries; southern South American islands; the central Kalahari and Sahara deserts; the Gobi Desert, Mongolia, and all of north-central and western China; from north of the Caspian Sea, Russia, to central Kazakhstan; and the “Outback” of central Australia. Five Arabian countries lack both samples and published records of indigenous diplopods – Bahrain, Kuwait, Oman, Qatar, and United Arab Emirates – as do Turks and Caicos, in the New World, and Mauritania and possibly Egypt, Africa. New records, including the first for Chilognatha from Botswana and the first specific localities from Northern Territory, Australia, are cited in the Appendix. Increased emphasis on mappings in taxonomic research is warranted along with investigations of insular “species swarms” that constitute a microcosm of the early evolution of the class. The largest “species swarm” in the Diplopoda is Diplopoda itself!
Characterized by small body size, apically rounded/lobed anterior gonopod telopodites, long slender posterior gonopod telopodites, and torsion in the cyphopod receptacles, Floridobolus fl oydi, n. sp., is described from the southern sector of the Brooksville Ridge in northwestern peninsular Florida. It inhabits sandy “Big Scrub” environments like F. penneri Causey, 1957, and F. orini Shelley, 2014, and is documented from the sector’s center and northern periphery, in Hernando and Citrus Counties, respectively, with a sight record from the eastern periphery. Its discovery supports the thesis that each sand ridge in peninsular Florida may harbor a unique species of this endemic genus.
With the discovery of Mitocybe auriportae Cook and Loomis, 1928 (Platydesmida: Andrognathidae) in Alameda County (Co.), east of San Francisco Bay, a potential overall distribution in coastal California is projected based on those of partly congruent diplopods. The area extends from northern Mendocino to central Monterey cos. and inland to central Lake, Yolo, and Santa Clara cos.